It became the major allele in wolves, making them bigger than other canids, probably during the last ice age. However, the small allele never disappeared.
During the #domestication time, humans created the first small dogs using the ancestral small allele that still existing in wolves. We found it in a 9,500 year old dog sample. Today this small allele is the major allele in modern dogs, in 75% of small breeds < 15kg